detect and identify
Bioluminescence
Bioluminescence
An overview by Manfred Hennecke,
BERTHOLD TECHNOLOGIES, Bad Wildbad, Germany
Bioluminescence is the ability of living organisms to
emit light. It is a phylogenetically widespread
phenomenon, that has evolved independently
many times in bacteria, fungi, marine plankton,
squids, fish, earthworms and beetles, but not in flowering plants and higher vertebrates (amphibians,
reptiles, birds and mammals).
The LU-LU reaction
Already 1647 Thomas Bartholin, a Danish physicist,
wrote a book on 'Animal Lights'. However, it
was only in 1885 that Raphael Dubois, a French
physiologist, demonstrated, that three substances
are involved in bioluminescence: luciferase,
luciferin and molecular oxygen. Luciferases (abbreviated as LU) are a class of enzymes that catalyze
the bioluminescence reactions. In most cases the
class of substrates are designated as luciferins
(also abbreviated as LU).
In general bioluminescent reactions or LU-LU
reactions can proceed with the above mentioned
minimum of three components and up to a maximum
of six components (luciferase or photoprotein,
luciferin, oxidant, cofactors, cations and a fluorophor
(a molecule which can absorb light of a given wavelength and emit light at a longer wavelength) to
generate living light.
Luciferases
American Firefly luciferase (Photinus pyralis, EC
1.13.12.7) is a 62 kDA protein, which is active as
a monomer and does not require subsequent
pro-cessing for its activity. The enzyme catalyzes
ATP-dependent D-luciferin oxidation by oxygen into
oxyluciferin with emission of light centered on 560
nm. As with many enzymes, firefly luciferase follows
Michaelis-Menten kinetics, and as a result maximum
light output is not achieved until the substrates and
co-factor are present in large excess. When assayed
under these conditions, light emitted from the
reaction is directly proportional to the number of
luciferase enzyme molecules.
Renilla luciferase, a monomeric 36 kDa protein,
catalyzes coelenterazine oxidation by oxygen
to produce light. The enzyme does not require
post-translational modification for its activity and
may function as a genetic reporter immediately
following translation. Coelenterazine native is the
natural substrate for Renilla luciferase. Meanwhile,
over a dozen of coelenterazine analogs have been
synthesized, which function as substrates for Renilla
luciferase with different properties in terms of
emission wavelength, cell membrane permeability
and quantum efficiency. Coelenterazine also emits
light from enzyme-independent oxidation, a
process known as autoluminescence.
In dinoflagellates, the luciferin is normally bound to
a protein, called luciferin binding protein (LBP). At
neutral pH, LBP stabilizes the luciferin from being
spontaneously oxidized. When activated by a
drop in pH, the luciferin dissociates from LBP, and
associates with a protein, luciferase, which acts as
a catalyst. In dinoflagellates, the wavelength of
maximum emission is approximately 470 nm, in the
blue-green region of the visible spectrum. In dinoflagellates Lingulodinium polyedrum, luciferase is
located in organelles called scintillons that emit
brief and bright flashes of light that are regulated
by an endogenous circadian clock. The complete
luciferase molecule has a molecular mass of 137
kDa and contains three homologous domains, each
of which is a separately active luciferase.
Bacterial luciferase from Vibrio harveyi is a heterodimer consisting of an alpha- (40 kDa) and a
beta-subunit (37 kDa), emitting light centered on
490 nm. Also luciferase from Photobacterium
fischerii (EC 1.14.14.3) is composed of two distinct
subunits, a and b, each of approximately 40 kDa.
The luciferase a-ß dimer has but one reduced flavin
binding site. pH optimum is 6.8.
Luciferins
In general, luciferin is an organic moiety that is
becoming oxidised in a bioluminescent reaction,
known as the bioluminescent substrate. Although
there are hundreds of types of luminous animals in
the sea, there are surprisingly few basic luciferins,
which have been found in across many species.
In some cases this conservation can be explained
by animals acquiring luciferin through the food
chain, but in other cases organisms have been
shown to have the ability to synthesize the same
chemical on their own. Today, there are five known
distinct chemical classes of luciferins, namely,
aldehydes, benzothiazoles, imidazolopyrazines,
tetrapyrroles and flavins.
Bacterial luciferin is
a reduced riboflavin
phosphate (FMNH2)
which is oxidized in
association with a
long-chain aldehyde,
oxygen and a luciferase. It is found in bacteria,
some fish and squid, which are hosting these luminescent bacteria in specific organells.
Physics - characteristics of the
light emission
Dinoflagellate luciferin
is thought to be derived from chlorophyll
and has a very similar
structure. In the genus
Gonyaulax, at pH 8 the
molecule is "protected"
from the luciferase by a
"luciferin-binding protein", but when the pH
lowers to around 6, the
free luciferin reacts and light is produced.
The emission of light resulting from the electronic
transition is the same process as fluorescent or phosphorescent light emission in-vitro. The end product of
a bioluminescent reaction is visible light recognized by
other organisms as a signal.
A modified form of this luciferin is also found in
euphausiid shrimp, perhaps indicating a dietary link
for the acquisition of luciferin.
Vargulin is found
in the ostracod
("seed shrimp")
Vargula, and is
also used by
the midshipman
fish Porichthys.
Here there is a clear dietary link, with fish losing
their ability to luminesce until they are fed with
luciferin-bearing food.
Coelenterazine is the
most "popular" of the
marine luciferins, found
in a variety of phyla.
This molecule can occur
in luciferin-luciferase
systems, and is famous
for being the light emitter of the photoprotein "aequorin". It is found in Radiolarians, Ctenophores,
Cnidarians, Squids, Copepods, Decapod Shrimps,
Mysid Shrimps, some Fish and Chaetognaths.
Firefly luciferin is used
in a luciferin-luciferase
system that requires
ATP as a cofactor.
Because of this, it can be used as a bioindicator of
the presence of energy or "life". It is found in fireflies and click beetles.
Other or unknown mechanisms are found in
Amphipods, Nemertean worms, Polychaete worms,
Bivalves,
Larvaceans,
Tunicates,
Limpets,
Earthworms and Fungus gnats.
During the chemical reaction in bioluminescence, light
is emitted as a result of generating an electronically
excited state. Electrons in excited orbitals decay to the
ground state and thus emit a photon, the light arising
from the potential energy of electronic transitions within
atoms or molecules.
The light emitted has very precise characteristics in
terms of color, intensity, polarization and timing.
Maximum emission at
Dinoflagellate
Renilla LU / Coelenterazine
Vibrio fischeri and harveyi
Photorhabdus luminescens
Earthworm LU / LU
Firefly LU / Luciferin
Click beetle
(Pyrophorus spec.)
470 nm
475 nm
490 nm
500 nm
500-550 nm
(species dependent)
560 nm
537 and 613nm
(Promega Notes 85)
Spectra of bioluminescence
lightblue:
darkblue:
Background:
firefly
Photorhabdus luminescens
Spectral sensitivity of a CCD camera
The geometry of light emission is also of importance
as light is directed from a particular part of an
organism in a specific direction for a particular
function. This is clearly seen in the point sources of
light emission in photophores, where the angle and
direction of light emission is very precise.
Anatomic distribution
The tissue distribution within organisms of the
components of bioluminescent reactions is quite
varied. The anatomic distribution of these components
gives clues as to the source of component synthesis,
storage, transport and the functional role of the
luminescence.
One key anatomic organ is the 'photophore' or 'light
producing organ' demonstrated in many luminous
fish and very vividly in cephalopods. Photophores
are normally made up of complex photogenic (light
emitting) cells.
Bioluminescent reaction components have also
been detected in the stomach, secretory organs
and liver of some organisms (mostly believed to be
there as a result of synthesis or storage).
Ecology and Behavior
As a result of its prevalence bioluminescence plays
an important role in the ecology of the oceans.
The function of bioluminescence in the oceans is
more clearly understood in the context of the
frequently dark environment. Some of the major
functions of bioluminescence are listed below:
Camouflage
Some marine animals that live near the surface
have luminescent organs on their underside
utilising bioluminescence in counterillumination
against the night sky. Bobtail squid e. g., when
viewed from underneath, disappears against the light
of the moon and stars. Or when disturbed, one species of squid emits a cloud of luminescent water
instead of the ink that its shallow-water relatives use.
Attraction
Bioluminescence is used as a lure to attract prey by
several deep sea fish such as the anglerfish.
A dangling appendage that extends from the head
of the fish attracts small animals to within striking
distance of the fish.
The cookie cutter shark is thought to utilize a
bioluminescent patch on its underbelly to appear as
a small fish to large predatory fish like tuna and
mackerel. When these fish try to catch the "small
fish", they are attacked by the shark.
Dinoflagellates have an interesting twist on this
mechanism. When a predator of plankton is sensed
through motion in the water, the dinoflagellate
luminesces. This in turn attracts even larger
predators, which will catch the would-be predator
of the dinoflagellate.
The attraction of mates is another proposed
mechanism of bioluminescent action. This is seen
actively in fireflies, who utilize periodic flashing
in their abdomens to attract mates in the mating
season. In the marine environment this has only
been well-documented in certain small crustacean
called ostracod. It has been suggested that pheromones may be used for long-distance communication, and bioluminescent used at close range to
"home in" on the target.
The honey mushroom attracts insects using
bioluminescence, the insects will help disseminate
the fungus spores into the environment.
Repulsion
Certain squid and small crustaceans utilize
bioluminescent chemical mixtures or bioluminescent
bacterial slurries in the same way as many squid
use ink. A cloud of luminescence is expulsed
confusing or repelling a potential predator while the
squid or crustacean escapes to safety.
Communication
Bioluminescence is thought to play a direct role in
communication between bacteria. It promotes the
symbiotic induction of bacteria into host species,
and may play a role in colony aggregation.
Several deep sea fish communicate by the blue
light of their light organs.
Geographic distribution
The occurrence of bioluminescence is highly varied
in geographical terms, but mostly in marine
organisms. Since the visible area of the spectrum is
400-700 nm, the emission maxima of most marine
species falls within the range of 450-490 nm.
Species found in the pelagic environment in the
oceans are mostly blue-emitting whereas terrestrial
organisms are predominantly yellow-green emitting.
In oceanic water blue-green (approximately
400-500 nm) luminescence achieves maximum
transmission. Interestingly, visual pigments of
most marine organisms are most sensitive in this
area. On land Brazil hosts the greatest diversity of
luminescent insects in the globe.
They are mostly fireflies, click beetles and railroad
worms among other groups. About 500 species
have been described, but a much greater number
remain to be described. North America is known for
its fireflies, with a variation in species as compared
to Southern Europe and Asia.
Several glow-worm colonies have been identified in
Europe. Interestingly, some species are luminous
in one location and not in another e.g. the
fish Porichthys notatus. The luminescence of one
population of this species has been postulated to
relate to the availability of luminous dietary sources
in a particular area. In addition, 'red tides,'
which are 'blooms' of luminescent phytoplankton
(dinoflagellates), are well known in Puerto Rico and
Jamaica. One location in Puerto Rico named the
'Bioluminescent Bay' is well known for spectacular
observations of dinoflagellate luminescence.
Around Japan the firefly squid (Watasenia) displays
spectacular luminescence and is found in large
numbers in restricted localities. Small crustaceans
such as ostracods have also been found in abundance in Japanese coastal waters and were used
for map-reading during war time. These small
organisms can be dried and then re-hydrated with
water to produce light.
Phyletic distribution
Some 17 phyla and at least 700 genera contain
luminous species. Many luminous organisms in
the deep sea still remain to be characterised in
terms of the chemistry of their bioluminescence
components. Luminescence has been demonstrated
in cephalopods, copepods, ostracods, amphipods,
euphausiids and many fish, annelids and jellies to
name but a few marine species. On land, fireflies,
glow-worms and click beetles are just a few of the
recognised luminous species.
The following
give an idea
organisms (it
and definitely
list on the next page is intended to
of the diversity of bioluminescent
is not meant to be comprehensive
not complete on species level).
Bioluminescence in
Marine Animals
Invertebrates
Many single-celled organisms are bioluminescent
when disturbed. One well-known area to see
luminescent dinoflagellates is in Puerto Ricos
Bioluminescent Bay, on the islands southwest
coast. The lagoon is attached through a narrow link
to the Caribbeans gentle tides and vitamin-rich
water. The water contains up to 200,000 singlecelled bioluminescent dinoflagellates Pyrodimium
bahamense per litre.
These organisms emit a flash of bluish light when
agitated at night e. g. by boat or even dipping the
hand in the water.
Other disturbance is caused e. g. by dolphins gliding.
Such flow-induced bioluminescence provides a unique
opportunity for visualizing the flow field around
a swimming dolphin. Laboratory experiments using
fully-developed pipe flow revealed that the bioluminescent organisms identified in the field studies
can be stimulated in both laminar and turbulent
flow when shear stress values exceeded a certain
amount.
The radiolarian Tuscaridium cygneum forms
colonies in the deep-sea and glows when disturbed.
Also the deep-sea scyphomedusa Atolla vanhoeffeni,
abundant throughout the world, produce an
incredible perpetuated luminescence display when
disturbed. Nudibranchs are not generally thought of
as bioluminescent organisms, but this pelagic form
Phyllirrhoe has the ability to produce light.
There are very few reports of the bathypelagic
ctenophore Bathyctena because it is typically found
deeper than 2000 meters. The yellow spots along
the body are sources of bioluminescence which are
released into the water when the animal is disturbed.
The ctenophore Bathocyroè is one of the most
abundant mesopelagic species, but because of its
fragility, it was only described in 1978, when it was
collected from a submersible. This genus, like the
siphonophore Bargmannia, can produce blue and
green luminescence. Another ctenophore Mnemiopsis leidyi - was found in the Black Sea.
Amphipholis squamata is a small polychromatic
hermaphroditic ophiuroid (brittle star; Echinodermata) distributed worldwide except in polar
regions. The species is luminescent and large interand intrapopulational variations were observed for
luminous capabilities. This variability, partially of
genetical origin, is associated with positive and/or
negative consequences on fitness.
Caecosagitta macrocephala is the only species of
chaetognath (arrow worm) which is bioluminescent.
The species is quite common at around 700 meters
depth in the Pacific and Atlantic Oceans, but its
luminescence was at first discovered in the 1990s.
The light is produced at the midpoint of the body by
large vacuolar cells on the edges of the anterior
fins. Ovoid or fusiform membrane-bound inclusions
within these cells are autofluorescent to varying
degrees, a trait that has been correlated with the
location of luminescent sources in many taxa. The
fluorescent subcellular bodies are likely sites
for storage of the components of the luminescent
reaction. Most contain a dense paracrystalline
matrix with small spherical inclusions. Others, even
within the same cell, lack internal organization,
possibly indicating organelles in which the luminescent compounds have reacted. Because luminescence is normally produced in conjunction with
an escape response, the cloud of light appears to
function as a diversionary display, a commonly
hypothesized role for expelled luminescence.
Bioluminescence of the medusa Periphylla is based
on the oxidation of coelenterazine catalyzed by
luciferase. Periphylla has two types of luciferase:
the soluble form luciferase L, which causes the
exumbrellar bioluminescence display of the medusa,
and the insoluble aggregated form, which is stored
as particulate material in the ovary, in an amount
over 100 times that of luciferase L.
Squid developed both strategies: hosting luminescent
bacteria and active bioluminescent based on the
oxidation of coelenterazine. One example for the
hosting is the symbiotic relationship between the
squid Euprymna scolopes and V. fischeri providing a
remarkable example of specific cooperativity during
the development and growth of both organisms.
For example, once the juvenile squid becomes
infected with the bacteria, maturation of the light
organ begins. Studies have shown that hatchling
squid fail to enlarge the pouches that become the
fully developed organ when raised in sterile seawater.
Thus, there is a direct consequence on the physical
maturation of the squid's light organ as a result
of its symbiotic relationship with V. fischeri.
Bacteria
Fungi
Vibrio harveyi
Vibrio fischeri
Photobacterium phosphoreum
Photobacterium leiognathi
Photorhabdus luminescens
Pleurotus lampas
Omphalia flavida
Mycena spec.
Lampteromyces japonicus
Armillaria mellea
Dinoflagellates
Pyrodimium bahamense
Radiolarians
Tuscaridium cygneum
Nematodes
Neoaplectana spec.
Steinernema spec.
Heterorhabditis spec.
Cnidaria
Scyphozoa
Hydrozoa
Anthozoa
Ctenophores
Nemerteanworms
Mollusca
Nudibranchs
Clams
Squid
Octopods
Limpet
Land Snail
Atolla vanhoeffeni
Aequorea victoria
Obelia spec.
Clytia noliformis
Renilla reniformis
Ptilosarcus gurneyi
Virgularia spec.
Osteocella septentrionalis
Ocyropsis spec.
Bathyctena spec.
Bathocyroë spec.
Beroa gracilis
Beroe forskalii
Mnemiopsis leidyi
1 species
Phyllirrhoe spec.
Pholas dactylus
Teuthida spec.
Colossal Squid
Mastigoteuthidae
Sepiolidae
Watasenia scintillans
Symplectoteuthis oualaniensis
Callistoctopus arakawai
Latia neritoides
Quantula striata
Annelid worms
MarineChaetopterus pergamentaceus
Polychaeteworms
Sylidfireworm
Eunice viridis
Earthworms
Pontodrillus bermudensis
Diplocardia longa
Diplocardia alba
Diplocardia eiseni
Diplotrema heteropora
Octochaetus multiporus
Spenceriella minor
Pycnogonids
Chaetognaths
Echinoderms
Sea stars
Brittle stars
Seacucumbers
Caecosagitta macrocephala
Amphipholis squamata
Hemichordateworms
Urochordates
Pyrosomes
Tunicate
Larvaceans
Pyrosoma tuberculata
1 Species
Centipedes
Millipedes
Crustaceans
Copepods
Ostracods
Amphipods
Decapodshrimp
Euphausiids
Cypridina hilgendorfii
Systellaspis debilis
Euphausia superba
Insects
Coleoptera
Lampyridae
Lamprocera spec.
Lampyris spec.
Luciola spec.
Microphotus spec.
Macrolampis spec.
Microdiphot spec.
Pyrogaster spec.
Pyropyga spec.
Elateridae
Phengodidae
Phrixothrix hiatus
Cenophengus spec.
Distremocephalus spec.
Diptera
Mycetophilidae
Orfelia fultoni
Arachnocampa spec.
Homoptera
Fulgora spec.
Collembola
Chordates
Sharks
Fish
cookie cutter shark
Diaphas spec.
Porichthys notatus
Hatchet Fish
Deep-Sea Angler Fish
Pinecone fish
Chauliodus macouni
Benttooth
Bristlemouth
Photoblepharon palpebratus
The luminescing bacteria are also advantageous to
the squid, a nocturnal forager, by erasing the
shadow that would normally be cast as the moon's
rays struck the squid from above, thus protecting
the squid from predators below.
The squid, in turn, provides a sheltering haven with
a stable source of nutrients for the bacteria. Again,
the cell density regulation of luminescence ensures
that the bacteria waste little energy on light
production until the high concentration of
autoinducer indicates that the cells have reached a
cell density high enough that the energy expended
on providing illumination for the squid is likely to be
well repaid in food and protection.
The example for active bioluminescence by itself
with Luciferase/coelenterazine is the deep sea
squid Watasenia scintillans. The squid has more
than 800 minute luminous organs distributed over
its ventral mantle and different clusters of
pigments. The ventral organs produce a steady
glow of light, whereas the arm organs emit flashes
of light at 470 nm. This bioluminescence will be
seen, when the squid comes inshore to lay
fertilized eggs.
Today researchers found luminescent pelagic octopuses, Amphitretus pelagicus. These animals are
about 30 cm is size and never touch the ground of
the ocean, even as larvae. Their bodys are translucent and some organs are luminescent.
Vertebrates (Fish)
Also fish developed both strategies: The light of the
flashlight fish, Photoblepharon palpebratus, is
produced by continuously-emitting luminescent
bacteria within the organs, but its display is
controlled by the fish. These animals, which live
along reefs in the Gulf of Elat, Israel, appear to use
their luminescent organs for such varied functions
as attracting prey, signaling other members of their
species and confusing potential predators.
Deep sea anglers are found at substantial oceanic
depths varying from 300 to 4,000 m. Any typical
bait on the end of a fishing line would no longer be
recognizable at these depths, so deep sea anglers
have a bioluminescent organ at the end of the line.
Millions of light producing bacteria cause the deep
sea angler's lure to light up. Only female deep sea
anglers have the lure and it is probably under her
control. The female deep sea angler wiggles its lure
from a long appendage on its forehead to attract its
prey. Some deep sea anglers, including a species of
the Netdevil anglerfish (Linophryne arborifer) have
a luminous structure with treelike branches
hanging from its chin.
Almost all marine bioluminescence is blue in color,
for two related reasons. First, blue-green light
(wavelength around 470 nm) transmits furthest in
water. The second reason for bioluminescence to be
blue is that most organisms are sensitive only to
blue light - they lack the visual pigments which can
absorb longer or shorter wavelengths. The ability to
produce and see red light, gives the Malacosteidae
family of fish a huge advantage in the deep sea.
Although the light doesn't travel very far, it lets
them see their prey, without alerting the prey or
any potentially curious predators.
To produce red light, the Malacosteidae use a
combination of filters and fluorescent material.
Light in the photophore (a light-producing organ)
doesn't start out deep red. Initially the light has a
short wavelength. This light is absorbed by a
fluorescent pigment inside the photophore, which
takes the energy and re-emits it as red light
(wavelength = 626 nm). Before it shines out into
the sea, the light is also filtered until it has a
wavelength of around 705 nm.
Because most fish do not have a visual pigment,
which is sensitive to red (705 nm) light, the
Malacosteidae must have an additional adaptation
to make them sensitive to the red light. In the
genus Aristostomias the fish bears an additional set
of photoreceptive pigments, which can pick up light
in the red region. Fish in the genus Malacosteus
show no photoreceptors, but developed antenna
pigments, a derivative of chlorophyll, which
function like a plant's chlorophyll.
The red light emitted by the fish is absorbed by a
special molecule, which acts like an antenna.
By capturing the energy in this way, this sensitizing
pigment can transfer the energy to the visual
pigments, which are usually only sensitive to bluegreen light.
All these remarkable adaptations highlight the
importance of bioluminescence in the interactions
among marine organisms and demonstrates the
value of in situ observations for understanding life
in the sea.
Bioluminescence in Terrestrial
Fungi and Animals
Some fungi can also emit light. Luminescent fungi
such as Armillaria mellea and Mycena spp. produce
a continuous (non-pulsing) light in their fruiting
bodies and mycelium.
It is believed that bioluminescent fungi use their
light to attract insects that will spread the fungal
spores, thus enhancing their reproduction. Some
nematodes are luminescent due to the presence of
symbiotic bacteria associated with them.
Nematodes of the genus Neoaplectana, Steinernema
and Heterorhabditis have a symbiotic association with
luminescent bacteria like Xenorhabdus luminescens and
thus exhibit luminescence. Some Centipedes, one
millipedes, one land snail, many earthworms and a
lot of beetles show bioluminescence in terrestrial
ecosystems.
Earthworms
Diplocardia longa is found on the Georgia coastal
plain in the sandy soil of lawns and at the edge
of pine forests. These earthworms exude a
luminescent slime, sticky and continuously glowing
involving coelomic fluid packaged in coelomocytes.
The earthworm luciferin is a simple aliphatic aldehyde, N-isovaleryl-3-amino-propanal.
Earthworm luciferase catalyzes the luminescent
degradation of the hydroperoxide adduct of
earthworm luciferin. However, comparative studies
indicate that the spectra of the bioluminescence of
different earthworms have different emission
maximum between 500 and 550 nm. Some data
supports species specific determinants of spectral
color that may represent an bioluminescence
resonance energy transfer system (BRET in nature).
Land snail
Quantula striata is regarded as the only land
gastropod in the world capable of true bioluminescence. This snail is normally collected in
secondary forest, lawns, rubbish dumps, under
concrete slabs and also in crevices along walkways
especially after rain.
The luminous organ is said to originate flashes of
yellow-green light beneath the mucous fold of the
head in juvenile stage, is known as the 'organ
of Haneda'. Another curious character is that
the mature snail sometimes loses the organ
and does not appear again. A social role for
luminescence probably suggested that the
aggregations of young snails are on finding
sources of food.
Insects
Among all taxonomic groups, the insects have
the largest number of luminescent species.
Luminescent species are found in Collembola
(springtails), Diptera (fungus-gnats; Mycetophilidae),
Homoptera and Coleoptera [Fireflies (Lampyridae),
click beetles (Elateridae) and railroad worms
(Phengodidae)].
Bioluminescence in insects assume different
biological functions: (sexual attraction) fireflies use
their flashes for courtship; (defence) click
beetles use their thoracic lanterns to startle
enemies; (illumination) railroad worms may use
their head lanterns to hunt their preys; (prey
attraction) fungus-gnat larvae and some click
beetle larvae that live in termite mounds use their
luminescence to attract preys.
Coleoptera
The order Coleoptera constitutes the largest
bioluminescent group in which several hundred
species are known to contain highly developed
photogenic organs. In most insects the light
produced is yellow-green as in Photinus and
Lampyris (Coleoptera). In larval and adult female
railroad worms, the light organs on the thorax and
abdomen produce green to orange light, while that
on the head produces red light.
The best understood luminous insects belong to the
families Lampyridae, Elateridae and Phengodidae.
The members of Lampyridae are called fireflies or
lightning bugs. Their immature forms are commonly
referred to as glowworms, while the adults are
called fireflies. Similarly, the individuals of
Elateridae are called wireworms or click beetles.
Scientists have discovered that the brightest
insect is the very large Pyrophorus noctilucus
(Elateridae), with a brightness of 45 millilamberts.
This insect is also known as the Jamaican
click beetle and the 'cucujo' beetle of the West
Indies. The immature forms of Phengodidae are
called railroad worms.
Lampyridae (fireflies)
The highest numbers of about 2000 firefly species
are found in warm, humid areas of the world,
especially in tropical Asia and Central and South
America. Some species, however, are found in very
arid regions of the world. In these arid regions,
larvae and adults can be readily found following
rains. All known firefly larvae have photic organs
and produce light.
The behavioral function of the larval light has
received considerable speculation and several
plausible theories have been proposed. However,
the most generally accepted hypothesis is firefly
larvae use their luminescence as a warning signal
(aposematism) that communicates to potential
predators that they taste bad because they have
defensive chemicals in their bodies.
These larvae also increase both the intensity
and frequency of their glow when disturbed.
An experimental study of whether mice could learn
to avoid glowing objects by associating a
larval-type glow with a bad tasting object further
supports the aposematism hypothesis.
Not all firefly species are bioluminescent as adults,
but of the species that are, one or both sexes use
a species specific flash pattern to attract a member
of the opposite sex. These bioluminescent signals
can take the form of anything from a continuous
glow, to discrete single flashes, to "flash-trains"
composed of multi-pulsed flashes.
Fireflies produce light via a chemical reaction
consisting of luciferin (the substrate) combined
with luciferase (the enzyme), ATP (adenosine
triphosphate) and oxygen. The way, how fireflies
turn their luminescent organs - called lanterns - on,
was discovered:
The luminescent cells of the lanterns are close to
cells at the end of the tracheoles (that bring oxygen to - and take carbon dioxide away from - the
insect's tissues).
These cells contain nitric oxide synthase (NOS),
the enzyme that liberates the gas nitric oxide
(NO) from arginine.
Nerve impulses activate the release of NO from
these cells.
The NO diffuses into the lantern cells and
inhibits cellular respiration in the mitochondria
(probably by blocking the action of cytochrome
coxidase)
With cellular respiration inhibited, the oxygen
content of the cells increases.
This turns on light production in the peroxisomes
that contain luciferase and luciferin-ATP (the ATP
is generated when the lanterns are dark). The quick
decay of NO probably contributes to the short
duration of the flash.
In most species of North American fireflies, during
a certain time of night, males fly about flashing
their species specific flash pattern. Females of the
same species tend to be perched on vegetation,
usually near the ground, and if a flashing male
catches a female's fancy, she will respond at a fixed
time delay after the last male's flash.
A short flash dialogue may ensue between the male
and female as the male locates her position and
descends to mate (McDermott 1958). The courtship
patterns of Japanese fireflies seem to show many
variations of this type of communication system,
as well as courtship behaviors that include
pheromones as well as photic signals (Ohba 1983).
It is generally assumed that most non-luminous
North American fireflies locate mates through the
use of pheromones.
Aspects of male flash patterns are also thought to
be affected by sexual selection. Female fireflies
have been shown to prefer certain characteristics of
a male's photic signal (such as increased flash rate)
and respond preferentially to males that possess
these "sexy" signal components.
Fireflies use their flashes to attract mates. The
pattern differs from species to species. In one
species, the females sometimes mimic the pattern
used by females of another species. When the
males of the second species respond to these
"femmes fatales", they are eaten! Recent evidence
also suggests that these female mimics are not
only acquiring food but also defensive chemicals
from their prey, which they themselves do not
produce in large quantities.
Other insects
Phengodid males in the tribe Mastinocerini (Brasilocerus,
Euryopa, Mastinocerus, Mastinomorphus, Phrixothrix,
Stenophrixothrix and Taxinomastinocerus) glow from
larval photic organs and are luminous throughout
their adult life. Like the female photic emissions,
these emissions appear to serve a defensive rather
than a courtship function.
A male Cenophengus ciceroi was observed glowing
from "two faint green spots, each lateral to the
midline in the last abdominal segments. These
spots glowed continuously and uncontrollably".
Males from the South American genus Pseudophengodes, have a large photic organ similar in
size and shape to those found in some fireflies.
These photic organs are not of larval origin and
appear to be used in pair formation in these few
species.
Through a modern phylogenetic analysis of the
cantharoid taxa (those include Phengodidae and
their closest relatives) not only hypothesize that
phengodids and fireflies are not each others closest
relatives, but that bioluminescence arose twice and
was lost once in this lineage of beetles. Phengodids
and fireflies (family Lampyridae) have traditionally
been thought to be the nearest relative of each
other mainly due to the fact both families are
bioluminescent.
Photic organs in Zarhipis, are present as bands
(at the base of the meso and metathroax and on all
but the last abdominal tergites,) or spots (on upper
lateral surfaces of abdominal segments one
through nine); photic emissions generally are
greenish-yellow.
The photic organs in Phrixothrix are composed of
two medial organs on the head (producing red
photic emissions) and 11 pairs of photic organs
(producing yellowish-green emissions) located
from second thoracic segment through the ninth
abdominal segment. The larvae are predacious and
feed on millipedes.
In Fulgora (Homoptera) the light organ is situated
only on the head. The light organs generally
originate from fat bodies, except in Arachnocampa
(Diptera) where these stem from the enlarged
distal ends of malpighian tubules. Light produced
by Arachnocampa is blue-green while that of
Fulgora is white. Thus, the colour of light emitted
varies with species and the variation may be
due to environmental factors or differences in the
structure of luciferase.
Impact of Molecular Biology and
Bioluminescence
Photoproteins
The cloning of various components of bioluminescent
systems has heralded major advances in biological
research. The calcium-dependent photoprotein
aequorin from the jellyfish Aequorea victoria
was cloned in 1985. Because the intensity of
luminescence from aequorin varies with calcium
concentration, it is now a well established method
of measuring of cell calcium in medical research.
interpretation of the experimental data by reducing
extraneous influences. The experimental and control luciferase enzymes used in the DualLuciferase® Reporter (DLR) Assay have distinct evolutionary origins. The firefly luciferase and the
Renilla (sea pansy) luciferase can discriminate between their respective bioluminescent substrates
and do not crossactivate.
Since the click beetle luciferase emits light at longer wavelength but using also luciferin as substrate, other test kits with two luciferases have been
developed.
In 1985 an ATP dependent firefly luciferase was
cloned. This luciferase, which detects and measures ATP, can be a good measure of food contamination. Living organisms obviously contain ATP. An
assay which detects ATP can therefore detect living
organisms. This is particularly useful for the
detection of spoilage. Many other luciferases have
been cloned including the sea pansy Renilla
reniformis luciferase and the South American click
beetle luciferase.
The Chroma-Luc™ Reporter vectors encode
luciferases that emit green and red luminescence.
Although the luciferases are 98% identical in amino
acid sequence, their peak luminescence wavelengths
are separated by >75 nm. Both colours of
luminescence are generated by a single addition
of reagent, which has been developed for
homogeneous assay of mammalian cells directly in
culture medium. These novel vectors are useful for
dual measurements where closely similar reporter
structures are preferred or where a single reagent
addition is desired.
Current Applications
of Bioluminescence
In last years very sensitive photo-multiplier tubes
and slow scan CCD cameras have been developed
enabling non-invasive studies of reporter gene
expression in living animals and plants.
Luciferases
From that variety of natural bioluminescence
sources firefly luciferase/luciferin, renilla luciferase/
coelenterazine and bacterial luciferase using
coexpressed fatty aldehyds as substrate have been
successfully transferred into research, diagnostic
and clinical applications.
For water quality/toxicity testing the luminous
marine organism Photobacterium phosphoreum is
used. When the organism is challenged by a toxin,
the respiration pathway is disrupted, resulting in a
decrease in luminescence.
A similar diagnostic test is widely used in the food
industry where contamination can be linked to light
emission as a result of using a test based on the
bioluminescent firefly luciferase. In order for the
luciferase to emit light, ATP is required (along with
added luciferin). If light emission is generated,
the presence of ATP is indicated and therefore by
implication contaminating bacteria. The DualLuciferase® Reporter (DLR) assay system contains
two different luciferase reporter enzymes that are
expressed simultaneously in each cell. Typically,
the experimental reporter is correlated with the
effect of specific experimental conditions, while the
activity of the co-transfected "control" reporter
gene provides an internal control, which serves as
the baseline response. Normalizing the experimental reporter gene to the activity of an internal
control minimizes the variability caused by differences in cell viability and transfection efficiency.
Thus, dual reporter assays allow more reliable
Summary
A wide range of recombinant proteins derived from
bioluminescent organisms now provide the basis of
major detection technologies in cell biology
and medical research. With better characterization
and understanding of this naturally occurring
phenomenon the application potential is extremely
significant in terms of disease detection and control
and also in our understanding of oceanic
ecosystems and forces which impact upon it.
Acknowledgements
Attached is a list of references and literature. The
article is without pictures of luminescent organisms. On different internet-websides beautiful
photographs of these organisms are shown, easy to
find with web-browsers.
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